ACTA PALAEOBOTANICA 27 (1): 9-26, 1987
K. SZCZEPANEK
LATE-GLACIAL AND HOLOCENE POLLEN DIAGRAMS FROM JASIEL IN THE LOW BESKID MTS. (THE CARPATHIANS)
P6inoglacjalny i holocenski profil pylkowy z Jasiela w Beskidzie Niskim (Karpaty)
ABSTRACT. Material obtained from the cores of peatbog sediments, ca 2 m in thickness,. taken in the Dukla Mts. (Low Beskids, Carpathians), was used for pollen analyses. Six selected levels were dated by the 14C method. These data are used to present the vegetational changes. of the peatbog surroundings since IO 300 B. P.
PRESENT-DAY NATURAL ENVIRONMENT
Characteristics of the region
The Low Beskid Mts. constitute the lowest and narrowest part of the Car pathian arch (Klimaszewski 1935; Starke! 1972). Tbey are ca 1830 km' in area and ca 100 km long, whereas the greatest width is up to ca 30 km. The boundary between the Western and Eastern Carpathians runs in this region. The system of mountain ridges extends as a rule in a SE-NW direction. The region is characterized by a very complex mosaic of orographic forms. The highest peak, situated close to the western boundary of the area, rises to an altitude of 999 m. Except for one other peak, which reacbes 996 m, no mountain top attains 900 m. The highest mountains are grouped at the opposite ends of the range, which extends in the direction of the parallel of latitude. The lowest floors of the valleys lie at an altitude of 300 m. Out of the numerous ranges of the Low Beskids, the Dukla Mts., lying in the central part of the region, are tbe lowest. Here the culminations of i•idges and hills reach altitudes from 520 to 720 m. The Dukla Pass (502 m a.s.l.), which is the lowest in the whole Carpathian arch, is situated in this area. The southern summits mark the European wate... shed, separating the catchment area of the Baltic Sea from that of the Black Sea.
~ ( lllJTT llt.&Wi1t;1 tt \ ' 10
The geological structure of the Low Beskid Mts. is also very complex. They are built of rocks belonging to the Carpathii'!,n flysch, Lower Cretaceous-Oligo cene in age. Five main stratigraphic-tectonic units have been distinguished here: the Magura nappe, Dukla folds, Silesian nappe, sub-Silesian nappe and the Skolska nappe. The morphology of the region reflects the basic stratigraphic -tectonic elements and the varied resistance of rocks to weathering. The higher parts of mountains are mostly of the Magura series and Dukla folds. These rocks are the poorest in alkali compounds and calcium carbonate. As regards mor phology, the rocks richer in calcium carbonate are characterized by a lower and gentler relief. The rocky substratum is strongly folded and jointed throughout the Low Beskid Mts. (Swidzinski 1953; Starke! 1972). In the SE-NW elevations both inclination and degree of scaliness of the main rock formations increase regularly. As a result, the northern and western slopes are generally more humid (and wooded) than the southern and eastern slopes. The Quaternary covers consist chiefly of loamy-stony or clayeystone col luvial and solifluctional sediments, 10-30 m tli~ck, overlying tbe slopes. The valleys are filled with fluvial sediments. Diversified acid and leached brown loamy soils, occupy the greater part of the L(}W Beskid area. In the mountain ranges they are usually of a medium thickness (up to 50 cn:1) with a consideru.ble skeleton content (especially where more resistant sandstones occur in the substratum). Soils of this type, occurring on slopes and notably under crops, are exposed to intense processes of deflation. Bro"' n soils, reddish-brown soils, pai·arendzinas, podsols proper, initial hydro genic soils and others appear more rarely. The leading factor that effects the climatic conditions in the Low Beskid 1\fts. is the relief. The intensity of the mountain climate increa.ses from the piedmont regions towards the highest ranges. In accordance with the classification of the climate of the Polish Western Carpathians (Hess 1965), two climatic zones can be distinguished in this area: a moderately warm zone, with mean tempera tures, calculated for many years, lying between 6 and 8°C, and a moderately cool zone with mean temperatures from 4 to 6°C. In this region the boundary between these two zones runs at an altitude of 500-570 rn. The zones represent a pluvial type of climate and the whole area belongs to the Carpathian climatic province (Guminski 1950). Fairly big differences in mean annual rainfall from ca 700 to ca 900 mm occurring between particular areas are dependent on the relief and altitude. The region under discussion is characterized by the occurrence of strong southern foehn-typ€ winds blowing from over the Hungarian Lowland. These winds are most frequent in late autumn, ia winter, and in early spring, occurring more rarely in summer. They bring about great anomalies in the prevailing weatlier. In the Low Beskid Mts. the growing season lasts from 215 days in the valleys to 182 days on the summits (Hess 1965, Hess et al. 1976). 11
The flora. of the Low Beskid Mts. is relatively poor. This is due to their small and weakly differentiated altitudes, and the uniformity of the geologic substratum. At present woods cover 35-50% of the total area here (Swi~s 1980, 1982). The most intensely wooded parts are the highest mountain ridges at the opposite ends of the Low Beskid w11ge. l\fosotrophic communities prevail in its plant cover. The forest communities show the most distinct differentiation according to altitude. Two vegetational zones have been distinguished here: the foothills zone and lower montane forest zone. Remnants of the hornbeam forest (Tilio-Carpinetum) occur in the foothill zone, whereas the beech forests (Dentario glandulosae-Fagetum), not very widespread acidophilous fir forests, beech forests with V accini1im myrtillus and fragments of other forest communities make up tbe lower montane forest zone. Alder woods (Alnetum incanae) predominate in the river and stream valleys. Both self-sown and planted woods and thickets composed of Alnits incana, Pinus sylvestris and Juniperus communis occur generally on the now uncultivated ground. Apart from the natural meadow communities of the orders Arrhena.theretalia and Molinietalia, large are::i.s are occupied by semi-uatural meadows and sy nanthropic communities undefinable phytosociologically (Sw i~s 1980, 1982; Grodzinsk!a 1968; Grodzinska & Pancer-Kotejo"' a 1965). The nearly co1nplete disappearance of the western Carpathian species and the so mewhat slower disappearance of the eastern Carpathian species are a charac teristic botanical feature of this area. In the botanical division of Poland, this mountain range is distinguished as a separate sub-region and with respect to flora it is intermediate between the west and east Carpathians (Pawlowski 1972). The beginnings of the colonization of the Low Beskids by fairly numerous groups of people from the circle of the Corded Ware Culture took place as late as the Younger Neolithic. The popufation increased in the Bronze Age, at the time of the development of the Protoslav Lusatian Culture (Sulimirski 1957, 1959; Machnik 1960, 1962; Zaki 1955). Colonization occurred chiefly in dells and larger river valleys and included communities whose husbandry was mainly based on cattle breeding and pasturage. The region became densely populated as late as the early a,nd late Middle Ages, or in the 13th-15th cc. On the turns of the 14th and 15th century the nomadic-pastoral and pastoral-agricultural population contributed to the devastation of the natura1 forests in nearly all convenient places in the mountains. This was responsible for marked changes in the local water and microclimatic conditions, increased floods, the deepening of river channels and valleys, drying of slopes, etc. Owing to the considerable depopulation of this area after 1947, fields, meadows and pastures have been turned into swamps, and succession mainly of Alnus incana,, Betula verrucosa and semi-natural herb communities have developed. 12
Des~ription of the• site
The material used for the palynological study comes from a peatbog situated about 20 km to the south-east of Dukla, in the south-eastern part of the Dukla Mts. (49°22'22" N, 21° 53'13" E), close to the state frontier. The nearest villages, Jasiel and Rudawka Jasliska, are at 9J distance of 3-5 km. The peatbog is 10 hectares in area ~md lies at an altitude of about 670-680 m; it is below the summit of a hill o.a its side slopiug at about 5° northeastnorth ward. It is in the headwaters of a small stream - a left-bank tributary of the Jasiolka (Fig. 1). The peat layer is on the average 1 m thick, the maximum thickness
Fig. I. 1\fap showing the location of .Jasiel mire of the organogenic sediment being about 2 m. The pea.tbog was d.rained by several ditches. On its surface there gi.·ow clumps of Salix sp. div., Alnus gluti nosa, A. incana, Betula verrucosa, Frangula alnus, J uniperus commitnis, V acci nium vitis-idaea, V. myrtillus and above all Phragmites communis and .21:folinia coerulea, which form large groups. Mosses of the genera Sphagnum and Poly trichum overgrow small areas forming a mosaic. Degraded meadow (pasture) communities on a minei·al substratum extend south and west of the peatbog. In the north and east a dense fir-beech forest comes near the edge of the peatbog. In the north the peatbog is drained by several streamlets which dissect the pro jecting rocky bank supporting the sediment layers lying at the bottom of the peatbog. The peatbog is situated within tbe lower montane forest zone with 13 beech and fir-beech forests now prevailing. The climatic conditions are charac terized by the following data: mean a.nnual temperature - ca 5.5°0, mean daily temperature - 5°0, mean annual maximum temperature - 9.7°0, me1n annual minimum temperature - 2.3°0, °\'egetational season - ca 200 days (Hess et al. 1976 ).
PALAEOECOLOGICAL STUDY
Methods
A series of sounding borings were made in the peatbog to determine the thickness of the organogenic sediments. A maximum organogenic sediment thickness of ca 2 m was found over an area of several bund!'ed square metres about 60 m away from the northern edge of the peatbog and west of the main drainage ditch. Two cores were take a, using the Russian borer 5 cm in diameter, at a distance of 60 m from the northern edge of the peatbog a.nd tl>e forest and about 10 m west of the drainage ditch, in 1972 and 1981. One core was obtained in 1972. In 1981 three borings were performed 0.5 m apart in more or less the same place, and 5 segments, 0.5 m long, were taken from each of them. The results of palynological analyses of the eore from 1972 are presented in Fig. 4. From one ofcores collected in 1981, samples, 1 cm3 in °\'olume, we're taken at 5 cm intervals for palynological analysis, and the .remaining material was secured in 5-cm segments for 140 dating. Tbe material from borehole 2 was divided into sections 5 cm long and washed in order to obtain macroscopic plant remains. The core of borehole 3 was WJ_'apped in plastic film and stored. No determinable fruits, seeds and leaves wei·e found in sediment samples examined for macroscopic plant remain content. The 1-cm3 sediments samples ( 40 samples from the borehole of 1972, Fig. 4, and 50 from the borehole of 1981, l!"'ig. 2, 3) were treated by Erdtman's acetolysis, two sta11dard pollen pellets of Lycopodium being added in the core of 1981 (Stockmarr 1971). Samples with mineral con tent were treated with HF prior to acetolysis. At least 400 pollen grains of trees and shrubs and the remaining sporomorphes were counted and identified. The calculation of percentage values was based on the pollen sum of trees. shrubs, dwarf shrubs, and herbs (P). The sporomorphs of aquatic, swamp and cryptogamous plants were excluded. The basis of the calculation of pereentage values for the sporomorphs excluded from the sum of pollen was the sum of all the sporomorphs (without the spores of Lycopodium and Sphagnum, Fig. 2). The AP sum was used as the basis for the calcubtion of percentage values for the core of 1972 aud AP +N.AP (without Sphagnum) for all the other sporo morphs. The {loncentration of sporomorphs was calculated by the method de scribed by Stockmarr (1971) only for the core of 1981. 14
Aiter the completion of pollen analysis 6 levels were selected in the sediment core of 1981 for uc dating. This examination was carried out in the Laboratory of the Institute of Physics, Silesian Technical University at Gliwioe, iu. 1985. The following age determination were obtained (uncorrected vears B. P.).: Gd - 2330 Jasiel 7 /35 cm 600 ±60 Gd - 1945 .Jasiel 20/100 cm 4570±50 Gd - 2332 Jasiel 23/115 cm 4960 ±90 Gd - 2333 Jasiel 31/155 cm 9270±116 Gd - 1847 Jasiel 36/180 cm 9880±120 Gd - 1846 Jasiel 43/215 cm 10340±110 There are no reservations concerning ·these results.
Stratigraphy of sediments
.A simplified Troels-Smith's (1955) system is used in the description. The sequence of fayers in the profile collected in 1981 is as follows: 0-20 cm peat, light-brown, moderately decayed, passing into living turf of Sphagnum peat at top, wjth many pieces of plant tissues (including small wood fragments). Tha4, DI+ 20-118 cm peat, black-brown, heavily decayed, with many pieces of plant t.issues (Phragmites and other Gramineae, rootlets of Carex, small pieces of wood), somewhat loamed at floor; Ph34, .Ag+ 118-142 cm clay, light-brown wheu freshly taken, dark-grey when dried, hard, with many pieces of plant tissues (chiefly Phragmites) . .As 3, Th3l, .Ag+ 142-152.5 cm peat, black-bi·own, heavily decayed, with many pieces of plant tissues (mainly Phragmites and other Gramineae, rootlets of Carex), compact when dry, somewhat loamed. Th34, .As+++, .Ag+ 152.5-161 cm clay, dark-grey, compact, shrinking little when dry, with many interlayers of plant tissues . .As 3, Th31, .Ag+ 161-188 cm peat with gyttja, dark-brown, heavily decayed, compact when dry, with mauy pieces of tissues of Phragmites and other Gramineae, rootlets of Carex; fairly large wood piece of Salix sp. at 170-175 cm. Th31-2, Ld + -2, .Ag+, .As+, Sh 1 IV 188-197.5 cm peat with gyttja, dark-brown, lieavily decayed, matted, with many pieces of plant tissues and rootlets and with flat crumbles of flysch rock, up to ca 1 cm; somewhat loamed. Th32, Ld 2, .As+ +, Gg +, .Ag+
' 15
197.5-217 cm peat wit.h a small admixture of gyttja, dark-brown, beavily decayed, strongly matted, little resilient, with ma,ny pieces of Phragmites and other Gramineae and with rootlets of Carex (some rootlets of cf. Carex gracilis), loamed. . Th32-3, Sh 2, As+, Ld 1, Ag+ 217-245 om clay, grey-brown, compact when The division of the pollen diagram into local pollen zones and their description The distinction of the local pollen zones was based on the percentage Pinus - Gramineae local pollen zone Pinus (min. 14.57 %, max. 28.24 %), Gramineae (min. 21.83 %, abs. max 59.81 %) and Cyperaceae (abs. max. 37.23 %) are dominant elements in the pollen spectra of this layer. Pinus type haploxylon, Larix, Juniperits and Picea are their constant components, but their pollen occur in very low percen tages and do not form continuous curves. The Alnus pollen appears spora dically. Betula and Salix pollen form rising continuous curves, and within the herb pollen, Filipendula shows a similar tendency. The occurrence of Pinus cembra in situ is confirmed by the presence of a big piece of wood at a depth of 230 cm. The mineral sediment in the root portion of this layer was marked by a very low pollen frequency so the calculation of the spectra was relinquished at two levels. A radiocarbon date of 10340±110 years B. P. was obtained for the sample at a depth of 215 cm, i.e. at tbe floor of the organogenic sediments. Salix -· Betula - Artemisia local pollen zone The lower boundary of this zone is marked by a tendency for some curves, abo"Ve all, those of Pinus, Betula, Salix, Artemisia and Chenopodiaceae to rise. The Pinus, Betula and Salix concentration curves show analogous tendencies. The absolute maximum percentage values of Salix - 30.5 %, Artemisia 16 - 9. 79 % , Chenopodiaceae -1.51 % and Ran1mculaceae -13.41% 1 are a charac teristic feature of this zone and so is the first culmination of Betula -13.75 % . Pinus type haploa;ylon, Larix, Juniperus, Picea and Alnits a.re still present in the spectra. Ulmus, Corylus and Alnus tend to form continuous curves; parti .cula'rly in the top samples. Gramineae pollen curve decreases consi derably (9.81 %). The floor part of this layer was 140 dated at 9880±120 B. P. and the root part at 9270±160 B. P. Pin1ts - Betula - Polypod?°aceae local pollen zone The lower boundary is marked by the maximum value of Pinus, the rise of Betula and decline of Salix curve and the rising Picea, Alnus, Ulmus and Corylus curves. In this zone the percentage values of Pinus (30. 7 %), Betula (17 .06 %), Picea (4.59 %), Filipendula (33.42 %) and Polypodiaceae (19.39 %) pollen achieve their absolute maxima. The Ulmus and Corylus curves show a very distinct tendency to rise. The continuom but low Quercns and Tilia curves make their beginnings. The Cyperaceae reach an absolute minimum (0.45 %). The Gramineae curve tends again to rise. The general pollen concen tration increases distinctly, notably Jn the case of such taxa as Betula, Ulmus a.od Corylus. Corylus - Ulmus local pollen zone The lower boundary is marked by a distinct fall in the Pinus, Betula, Salix and Picea pollen curves and rise in the Corylus, Alnus and Quercus curves. The zooe is characterized by the absolute maximum of Corylus (27 .65 %), relatively high percentage values of Ulmus (6.33%, 7.17%) and high concen tration of Pinus pollen, (its percentage values tending however to decrease) . .A rapid rise in concentratoin for Ulmus, Corylus, Querctts and Tilia pollen is very characteristic. As regards the herbs, the concentration of the Cyperaceae, 0 Gramineae and Filipendula pollen and Polypodiaceae spores also increases very much. Ulmits - Quercus - Tilia local pollen zone The further downward tendency observed in the percentage curves of Pinus a.i1d Betiila pollen and a distinct fall in Corylus mark the lower boundary of this zone. There is Po rise in the Ulmus curve and also a gentle but systematic rise in the Quercus curve. This zone is charactedzed by the absolute m3Jxima of percentage values for Ulmus (10.60 %), Querci1.s ( 4.96 %), 'l1Uia (2.32 %) and Fraxinus (2.32 %), the percentage of Corylus being still relatively high (13.5-17 .2 %). The course of the percentage curves of these taxa is fairly even, without major fluctuations. 17 There is a distinct upward trend in the Alnus curve (.max. 4.54 %). The presence of Pagus is represented by a continuous low curve and so is that of Oarpinus in the top portion. The maximum pollen concentrntion is also a characteristic feature of the zone. Pinits, Betula, Til-ia, Gramineae; Fiiipendu~a and Artemisia pollen achieve their maximum concentrations for the whole profile. Two 140 d~tes were obtained for this layer: 4960±90 years B. P. for the floor sample and 4570±50 years B. P. for the top one. Quercus - Carpinits local pollen zone The distinct rise of the percentage pollen curves for Alnvs, JPa,gits and Oar pinus, the beginning of the continuous Abies curve and a pronounced tendency of Fraxinus curve to decline form the lower boundary of this zone. The systematic though gentle decrease of Ulmus, Corylits and Tilia curves and the very low values of Pinus are characteristic features of this zone. The Fagus cur>-e rises and so does the curve for Abies but far more gently. Carpinius has an absolute maximum ( 4.37 %) here. The course of the Alnus curve is fairly even and the Gramineae curve rises distinctly. The cur~e of general pollen oon ct:lntration shows a continuous though mild downward tendency with the in creasing concent1 ations of Fagus and Garpinus pollen. Fagus - 11bies local polien zone The lower boundary is marked by the rise of Fagus curve to the maximum values and by a clear growth tendency in the Abies a:nd Sphagnum cur ves. The root portion of the sediments was 140 dated at 600±60 years B. P. Tbis zone is bipartite: the Fagus - Abies subzone and the .Alnus - Sphagnum subzone can be distinguished. The boundary between them is placed of a distinct rise in the Alnus and Sphagnum percentage values and the absolute maximum of Abies. Fagus - Abies subzone This subzone is characterized by the absolute maximum of Fagus (18.49 %), whose percentage curve has a regular even course. The Oarpinus ourve does not show any major fluctuations, as well. The Abies and Sphagniim curves rise systematically towa;rds their maximum values. Alwus (max. 5. 71 %) and Quercus (max. 3.81 %) curves have an even course at a relatively higl> level. The Ulmus and Corylus curves deoUne gently and systematically. The curves of Plantago lanceolata, Rumex and Secale rise . .A very slight rise in the t>ercentage pollen values of Artemisia (1.11 %) and Ghenopodiaceae (0.30 %) can be observed too. 2 - Acta Palaeobotanica 27/1 18 Alnus - Sphagnum subzone In this subzone the Abies curve reaches its absolute maximum (7 .24 %)r the Sphagnum curve shows relatively high values (ca 10 %) and forms a distinct peak (15.67 %) in the middle part of this layer. The rise of Alnus pollen curve (max. 34.5 %), with a tendency towards an absolute rnaximurn, is characterjstic. The pollen of synanthropic plants form already continuous though low curves. A more or less regular mean pollen concentration is characteristic of the whole of this local pollen zone except for one level at which it is somewhat higher. Alnus - · Secale local pollen zone The lower boundary of this zone is fixed on the basis of the nmximum rise of the Atniis curvre and the steep fall of the Fagus, Carpinus, Abies and Qttercus. This zone is characterized, above all, by the absolute maximum of Alnus (75.68%), accompanied by a marked fall in the Gramineae percentage curve (abs. min. 7.45 %). The Betnla curve shows a, slight upward tendency. The Fagus· and, to a much lower degree, Quercus and Corylus curves, after a preceding fall, also show a rising trend, and so do the Gramineae and Sphagnum curves. The· fall of the percentage pollen curves of nearly all taxa is here evidently caused by the over representation of Alnus pollen. The most distinct upward tendencies in this zone characterize the percentage curves of Plantago lanceolata (max. 1.41 %), Rumex (max. 0.50%) and Secale (max. 0.64%), although their percen tage values are still very low. The concentration curve descends regularly~. HISTORY OF CHANGES IN NATURAL ENVIRONMENT History of vegetation in the region of the peatbog at Jasiel during the last 10300 years· The reconstruction of vegetational changes in the surroundings of peatbog at Jasiel (Low BeskidMts., Carpathians) is based on the pollen anaJytical results obtained from two profiles and the radiocarbon dates of six levels of the profile collected in 1981. The composition of pollen spectra from t;he floor layers of sediments makes it possible to define the vegetation that accompanied the deposition of these layers as the vegetation of open areas. This is indicated especially by the AP NAP percentage ratio (AP forms less than 40 %). A fragment of Pinus cembra wood found at the floor of the organogenic sediments indicates its growing in situ. Rather low percentage values of pollen of Pinus cembra, Larix, Juniperus, Picea as well as Pinus type sylvestris, Betula and Salix indicate that these trees and shrubs have already played a certain role in the landscape. Representauts. 19 of Gramineae (tisstle of Phragmites), Cyperaceae (rootlets of Carex sp. and of Garex cf. gracilis were found), Rosaceae Ranimculaceae, Compositae, Umbelli Jerae families and Filipenditla itlmaria were dominant herbs, recording, the presence of swamps communities mainly . .A relatively small extent of xeric communities can be inferred from the not very high percentage -values of Arte misia and Chenopodiaceae and the lack of other indicators of this type of bio topes. Both, the 140 date of 10340±110 B. P. for the bottom portion of the orga nogenic sediments in the middle pa.ct of the Pinus-Gramineae local pollen zone and the general character of vegetation are indicative of the closing of time late glacial (Younger Dryas). The next stage of the vegetation development (Salix-Betula-Artemisia local pollen .zone) still has many traits of the previous period, that is, some contri bution of Pinus haploxylon, Larix, Juniperus, Picea and Alnits pollen to the spectra. The organogenic sediments contain a relatively large admixture of inorganic material, and alternate with mineral layers, what suggests a humid climate with high rainfall and deflation processes, removing mineral material from the slope. This stage is characterized by a distinct tendency for the forest communities to close. In the .AP to NAP ratio .AP is do:m.inant (from 45 to above 60%). It is chiefly composed of Pinus type sylvestris, Betula and Salix pollen. The expansion and closing of these trees are also confirmed by their pollen concentration curves, although the cumulative concentration ourve shows no significant changes as yet . .A piece of Salix wood at a depth of 170 cm is a direct evidence of its presence in situ. The fluctuations of Cyperaceae, Gramineae and Filipendula pollen values may have been connected with the fluctuations of the ground-water level. The maximum values of Artemisia and Chenopodiaceae pollen indicate an increasing role of plant communities in rather dry habitats. It is hard to interpret the high values of Ranunculaceae (13.41 %) in that period in respect of their ecological significance, for it may represent the species of both dry and very damp biotopes. Two 140 dates mark the time limits of the phenomena discussed above . .A sediment sample from the bottom layer was dated at 9880±120 years B. P. and that from the top layer at 9270±160 years B. P. Just above this date but still within the limits of the Salix-Betula-Artemisia local pollen zone the con tinuous uzm,us, Corylus and Alnus pollen percentage curves start i·ising, and their concentration curves rise also, evidencing the entering of these trees into the communities of the nearby surroundings. The two earlier stages of vegetation development in the surroundings of the peatbog resulted in a considerable expansion of forest communities during Pinus-Betitla-Polypodiaceae local pollen zone. These communities, may still have been rather loose, which is evidenced by the Polypodiaceae curve (abs. max. 30.78%); no spores of Dryopteris thelypteris were found. The Pinus type sylvestris (abs. max. 30. 78 %) and Betula pollen percentages (abs. max. 17 .06 %) 2* 20 were high. Pollen of both these trees and Polypodiaceae spores show also high concentrations. In this stage of the formation and closing of forest communities the spruce (Picea) also played a,n impoi·tant role. Its curve reaches here a.n absolute maxi mum for the whole profile (4.59%). Although theie ai·e no direct evidence, its history in the Carpathians (Srodo:ri 1967) and especially in the neighbouring Bieszczady Mts. (RalskD1-Jasiewiczow~ 1980) permits to suppose that the spruce grew in the Low Beskid lVHs. mainly in damp and peaty habitats. Ulmus, rising from 0.12 to 6.79% and, to a smaller degree, Corylus also become important components of the forest communities what is confirmed by the pollen concentration curves. Alnits shows no expansive tendencies yet, and Quercus appears in the area (the beginning of its continuous curve). The dominant trees and shrubs, forming dense forests, noticeably reduced the role of Salix. Changes occurring in the herbaceous vegetation consist chiefly of the re duction of swamps communities and of the open communities of heliophilous plants. These changes are reflected by the impoverishment of floristic com position of pollen spectra. The deposition of a loam layer shown in the profile of sediments in this phase suggests a period of increased rainfall (Starke! 1960, 1977; Gil et al. 1974). The following stage of the vegetation development ( Oorylus- Ulmits local pollen zone) brings an essential change in the composition of forests. Pinus and Betula are already of minor importance. The dominant species are CoryluB (abs. max. 27.18 and 27.65%) and Ulmus (6.33- 7.17%). Quercus, more slowly .Alnus (0.60-2.56%) and TUia (1.15%) grow in importance. Fraxinits appears in the area and the first pollen grains of Fagus are signs of its approaching. In the herbaceous communities the Gramineae (24.32 %) and Oyperaceae (6.27 %) occur in the largest numbers. The Polypodiaceae do not play a major role any longer and the frequency of Filipendula falls markedly (23-13 % ). The aquatic plants (Potamogeton) arc present in small quantities. In spite of a l:nge admix ture of organic matter (pieces of Phragmites tissues) the sediment is here mineral (loam). The pollen concentration in the sediment is very high. The -percentage values of Pinus and Betula are low but their pollen concentration is high. The concentration of Querciis and Tilia increases distinctly and that of Alnus at a considerably slower rate. Further changes in the plant cover ( Ulmus-Quercus-Tilia local pollen zone) go towards the growing importance of mixed deciduous forests composed of Ulmus (10.60%), Quercus (4.96%), Tilia (2.32%) andFraxinits (2.53%). Corylus (13-17%) plays an important role in their unoerstorey. Fagus (0.20-0.73%) . .Acer and Carpinus (0.95 %) are new elements. In the herbaceous communities the Grarnineae and Cyperaceae again show a small rise. A distinct enrichment of the swamp communities (Potamogeton, Sparganium and Lysimachia) is visible. The general pollen concentration attains the highest values with the maxi~ 21 mum pollen concentration values of Pinus and Betitla ·while their percentage values approximate to the minimum ones. These changes in the plant cover take place within a time interval from 4960±90 to 4570±50 years B. P. The changes of vegetation occurring in the region of the Low Beskid Mts. after ca 4500 B. P. include a slowly proceeding decrease in the Oorylus and Ulmus role and the spread of Oarpinits (abs. max. 4.37%), Fagus (max. 8.19%) and A.hies (max. 0. 71 %). They enter mixed deciduous forests, in which Qitermts, Tilia, Fraxinus, Acer and Alnus keep pl:c.ying an important role. Gramineae pollen prevailing in NAP (::"bove 40%), represents mainly Phrag mites. The proportions of Filipendula and Lysimachia are also notioeable. The aquatic plants are represented by Potamogeton. The higher rate of sediment accumulation is proba,bly responsible for the decline in the general pollen frequency curve, although tho dominant elements are still characterized by high values. The formation of fir-beech and hornbeam forests is completed during the next phase (Fagus-Abies local pollen zone), fir and beech forming lower the montane forest zone and the hornbeam the zone of foothill forests. The first distinct traces of human activity, that is rising continuous curves of Plantago lanceolata, Riimex, cf. Cannabis and Urtica are another characteristic feature of this place. The Artemisia and Ohenopodiaceae pollen percentage values also rise in the spectrr•. The rapid increase of the Alnus curve should be ascribed to increasing human activity and the colonization of post-agricultural areas by A. incana, which nowadays often plays the role of a pioneer species in. such habi tats in the Carpathians (cf. Bieszczady Mts. -Ralska-Jasiewiczowa 1980). The destruction of hornbeam forests is illustrated by the decliae in Ulmus, Oorylus, Tilia, Fraxinus and Acer pollen curves and the fluctuation of the Quercus curve. The fir-beech and high situated hornbeam forests did not suffer any sei·ious harm still at that stage. Changes in anthropogenic indicators are the basis for the distinction of two local pollen sub-zones. The rising Sphagnum curve must be referred to the rising humidity of climate. The date of the top portion of this phase, 600±60 years B. P., fairly well corresponds with the historical phases of settlement from the 13th c. onwards. The last stage of changes in the plant cover of this area (Alnus-Secale local pollen zone) comprises fluctuations ia tbe tree and herb polle.a values, which are connected with the human activity within the now prevailing plant communities. The influence of :man's husbandry is i·eflected by the occurrence of pollen of cultivated and synanthropic plants. In the pollen diagTam under study the curves for these plants are very low (abs. max. Secale - 0.64%, Rumex - 0.50%, Plantago lanceolata -1.41 %), which probably results from the type of farming, in which pasturage and animal breeding played an important role. Under the conditions of the increasing expansion of this type of farming Fagus aind Alnus seem to be the stablest components of tbe plant rover. 22 LOCAL AND REGIONAL HISTORY OF THE VEGETATION OF THE LOW BESKID MTS. AT THE DECLINE OF THE LATE GLACIAL AND IN THE HOLOCENE The results of studies on the fossil floras of the late glacial and the Holocene have been, as yet, published from the following localities in the region of the lowest depression in the Carpathian arch: Roztold near Jaslo (Szafer & Jaro:ri 1935; Szafer 1948), Cergowa Gora near Dukla (Wi~ckowski & Szczepanek 1963), Besko near Sanok (Koperowa 1970). K~pa near Krosao (Gerlach et al. 1972) and Szymbark-Kamionka near Gorlice (Gil et al. 1972; Gil et al. 1974) . .As a contribution to the IGCP Project No 158 K. Harmata again carried out a palaeobotanic investigations of sediments in the region of Roztoki near Jaslo (this volume). Several layers of sediments from Besko, K~pa, Szymbark Kamionka and Roztoki were dated by the 14C method. There is a comprehensive monograph of the area bordering on the Low Beskid Mts. in the ea.st written by Ralska-Jasiewiczowa (1980), whereas in the west the nearest p£ofiles ana lysed are those of a peatbog at Bryjarka (Pawlikowa 1965) and somewhat more distant peatbogs in the Nowy Targ Basin (Koperowa 1962; Obido wicz MS). The sites in the Low Beskid Mts. and neighbouring Doly Jasielsko-Sanockie studied by the palaeobotanic methods are situated withia the limits of two adjoining climatic-vegetational zones of the Carpathians and provide the bases for the investigation of the history of vegetation from the decline of the late glacial throughout the Holocene. The occurrence of Pinus sylvestris, P. cembra, Betula type alba, B. nana, Larix, Juniperus and Salix, and of plants of dry habitats, also of importance here and represented chiefly by Artemisia and Chenopodiaceae, is a common feature of the plant cover of this region till the spread of trees and shrubs with greater thermal requirements. The presence of Pinus cembra evidenced in situ gives the plant cover of this area a montane character and distinguishes it from late ghcial floras from the lowlands. Pinits sylvestris played an import::mt role in the vegetation of the moun tainous parts of the Low Beskid Mts. at that time. It was the codominant or dominant tree in the plant cover . .As the deciduous trees appeared and expanded, Piniis sylvestris persisted in la.rger valleys, being a retreating species in higher situations. Picea abies is another interesting species in this area. Its macroscopic re mains found at the sites in Doly J asielsko-Sanockie (Besko, K~pa - needles, Roztoki - seeds) are accompanied by very low slight percentage values in the pollen profiles (Roztoki, Bosko, K~pa, J asiel). In the Low Beskid the occurrence of spruce is very distinctly attached to peatbogs. The spruce played a certain role in these habitats in the initial stages of the formation of Holocene forest communities. In the younger Holocene it did not form a separate vegetational zone here and its distribution was clearly conditioned by the lack of suitable 23 habitats. In the ranges of the East and West Carpathians, neighbouring on the Low Beskids, the spruce had a significant role in the forest conununities during the climatic optimum of the Holocene. It was only the expansion of Oarpinus and particularly of Fagus and Abies that reduced the area of its occurrence chiefly to the higher locations where it formed a separate vegetation zone .(upper montane forest zone). The occurrence of Alnus in the Doly Jasielsko-Sanockie region from the decline of the late glacial onwards is also confirmed by macroscopic remains (Roztoki - seeds) and, sometimes, by significant percentage values of pollen (Besko - 5%). As the formation of mixed deciduous forest proceeds, Alnus (probably A. glutinosa) is a forest-forming element in lower locations and valleys of the area studied. The marked restriction of Alnus role, as the grounds in the valleys deforested by man increase in area is a characteristic feature in its Holocene history. On the other hand, in the more humid climatic phases in the younger Holocene, Alnits (or more strictly A. incana) played a pioneer role in the colonization of the grounds situated on steeper and more humid slopes no longer cultivated by people. In the younger Holocene Abies, Oarpinus and Fagus are codominant ele ments of the forests of the Low Beskid Mts., just as they are in the other ranges of the Carpathians. In the pollen diagrams from sites in the Low Beskid region the continuous curves for Carpinus and Fagus begin nearly at the same time. It is the rule that Carpinus attains its maximum earlier, and later its frequencies decrease. The Fagus curves are, as a rule, much higher and indicate the essential role of this tree in the communities of the lower mountain forest zone in the young Holocene. The beginning of the continuous Carpinus and Fagus pollen curves piecedes somewhat the date of 4500 years B. P. and usually precedes also the first traces of human activity. In a majority of the localities the con tinuous Abies curve begins much later, and attains significant values what cor responds with the role of this tree in the forests. The continuous Abies curve seems to coincide of ten with the first appearance of the culture pollen indicators. Human hmbandry may have exerted a decisive influence upon the role of the hornbeam in the forest communities of lower situations, for these places were most convenient for settlement. In consequence, the percentage values of Carpinus pollen in pollen diagram were lower than these of Fagus or Abies, which occupied the higher lying areas, less suitable for farming. Starting from this Rtage, the increasingly large areas brought into cultivation by man as well as the type of forest exploitation had a great influence upon the vegetation. In the author's opinion however, human husbandry was not the decisive factor influencing the transformation of the plant communities in this region (Sro doi1 1985). The climatic changes constituted the primary factor .. The early appearance of Carpinus and Fagus, considerably preceding that of Abies and the very distinct dominance of Fagus indicate the closer connec tion of forest communities i.o. the Low Beskid with those of the East than those of the West Carpathians. Also because of the minor role of Picea in the younger 24 Holocene the vegetation of the Low Beskids more resembles the plant cover of the Bieszczady Mts. than that of the West Beskids. The herb curves in the pollen diagrams reflect clearly the local vegetation. The relatively low curves for synanthropic and cultivated plants in all the pollen diagram result not so much from the late penetration of these areas by human husbandry as rather from its pastoral-agricultural nature and the still existing large wooded areas (present forestation: ca 35-50% ). The noticeable differences in the sediment accumulation rate at some locali ties are possibly connected with the fluctuations of humidity. In the profile from Jl'Jsiel there are at least two distinct layers of loam separated by a several -centimetre-thick layer of peat. The age of the first of them is determined by the date of the floor sample of the overlying peat, i.e. 9270±116 yeu.rs B. P. The other loam layer is com prised between this date and that of 4960±90 years B. P. The second silty sediment layer, 35 cm thick, covers a time intervr,I of nearly 4000 years and includes 3 local pollen zones (Salix-Betula-Artemisia, Pinus-Betula-Poly podiaceae and Oorylus-Ulmus). The floo.r of this layer is palynologically refered to the Boreal period. The period following 8500 B. P. known in the Carpathians and their foreland as a period of active hydrological pr0cesses resulting in the arising of landslips (e.g. Szymbark-Kamionka), inundation of depressions (e.g. Besko, Roztoki), i.nterealations of flood mud in the sediments of peatbogs on river terraces e.g. Tarnawa in the Bieszczady Mts. (Ralska-Jasiewiczowa. 1980; Starke! 1975) . .A slower i·ate of sediment accumulation in pel1tbogs in this time interval can also be found outside the Carpathians (e.g. Slopiec in the Swi~tokrzyskie (Holy Cross) Mts. - Szczepanek 1982). In particular cases this slackening may have been connected with the movements of ground-water. In the mountains, especially on the slopes, we should also take into account the processes of solifluction which are closely associated with humid climatic phases. ACKNOWLEDGMENTS I wish to express my heartfelt thanks in the first place to Dr. R. Soja for suggesting th& peatbog and helping me with boring in the field, Assist. Prof. l\f. Ralska-Jasiewiczowa for valuable remarks and discussions, Ms Z. Tomczynska for the identification of wood samples, Ms D. Nalepka for the identification of plant tissues from several layers and Ms B. Nowaczynska for the preparation of samples for palynological analyses and another technical help. Institute of Botany, Botanical Garden, Jagellonian University, ul. Kopernika 27, 31-501 Krakow Instytut Botaniki Uniwersytetu Jagiellonskiego, Ogr6d Botaniczny REFERENCES Gerlach T., Koszarski L., Koperowa W. & Koster E. 1972. Sediments lacustres post glaciaires dans la depression de Jaslo-Sanok. Studia Geomorph. Carpat. - Balcan, 6i 37-61. 2.5· Gil E., Kotarba A. & Szczepanek K. 1972. The site Il-3 the landslide at Szymbark-Kamion ka. Exe. Guide-Book. INQUA Holocene Syrop. 1, Poland: 42-45. Gil E., Gilot E., Kotarba A., Starkel L. & Szczepanek K. 1974. An early Holocene land sliche in the Beskid Niski and its significance for palaeogeographical reconstructi~ns. Studia. Geomorph. Carpat. - Balcan, 8: 69-83. Grodzinska K. & Pancer-Kotejowa E. 1965. Zbiorowiska lesne Pasma Bukowicy w Bes kidzie Niskim. (summary: Forest communities of the Bukowica Range (Low Beskids,. Polish Western Carpathians). Fragm. Flor. Geobot., 11 (4): 563-599. Grodzinska K. 1968. Rosliny naczyniowe Pasma Bukowicy (Beskid Niski). (summary: The vascular plants of the Bukowica Range (Low Beskids, Polish ·western Carpathians). Fragm. Flor. Geobot., 14 (1): 3-82. Guminski R. 1950. Waznicjsze elcmenty klimatu rolniczego Polski Poludniowo-Wschodniej. Wiad. Slu:Zby Hydro!. Meteorol., 3 (1): 57-113. Harmata K. 1987. Late - Glacial and Holocene history of vegetation at Roztoki and Tar nowiec near Jaslo (Jaslo - Sanok Depression). Acta Palaeobot., 27 (1): Hess M. 1965. Pi~tra klimatyczne w polskich Karpatach Zachodnich. (summary: Vertical climatic zones in the Polish vVcstern Carpathians). Zesz. Nauk. UJ, 115, Prace Geogr., 11, Prace Inst. Geogr., 33: 1-267. Hess M., Niedzwiedz T. & Obr~bska-Starklowa B. 1976. Stosunki termiczne Beskidu Niskiego (metodyka charakterystyki rezimu termiczncgo g6r). Inst. Geogr. i Przestrzen nego Zagosp. PAN. Prac. Geogr., 123: 1-101. Klimaszewski M. 1935. Z fizjografii Beskidu Niskiego. Wicrchy, 13: 89-93. - 1946. Podzial morfologiczny Poludniowej Polski. Czas. Geogr., 17 (3-4): 133--182. Koperowa W. 1962. P6inoglacjalna i holocenska historia roslinnosci Kotliny Nowotarskiej (summary: The history of the Late Glacial and Holocene vegetation in Nowy Targ Basin). Acta Palaeobot., 2 (3): 1-57. 1970. P6inoglacjalna i holoce1iska historia roslinnosci wschodniej CZ~Sci Dol6w Jasielsko -Sanockich (summary: Late Glacial and Holocene history of the vegetation of the eastern· part of the "Jaslo-Sanok Doly" Flysch Carpathians). Acta Palaeobot. 11 (2): 1-42. Machnik J. 1960. Ze studi6w nad kultur~ ceramiki sznurowej w Karpatach Polskich (summary: From studies on the Corded-Ware Culture in the Polish Carpathians). Acta ArehaeoL Carp., 2 (1-2): 55--86. 1962. Uwagi o zwi~zkach i chronologii niekt6rych znalczisk kultury ceramiki sznurowej w Karpatach (resume: Observations sur les connezitis ct la chronologie de certaines trou vialles de la civilization de la ceramique corolee dans les Carpates). Acta Archaeol. Carp., 4 (1-2): 91-107. Obidowicz A. (MS). Postglaziale Vegetations, Klima- und Besidlungsgeschichte der West karpaten. Pawlikowa B. 1965. Materialy do postglacjalnej historii roslinnosci Karpat Zachodnich. Torfowisko na Bryjarce (summary: Materials for the Postglacial history of vegetation of the West Carpathians. Peat-bog on the Bryjarka). Folia Quatern., 18: 1-9. Pawlowski B. 1972. Szata roslinna G6r Polskich. In: Szafer W. Za.rzycki K. (eds.). Szata Roslinna Polski, 2. PWN, Warszawa. Ralska-Jasiewiczowa M. & Starke! L. 1975. The leading problems of palaeogeography at the Holocene in the Polish Carpathians. Biul. Geolog., 19: 27-44. Ralska-J asiewiczowa M. 1980. Late-Glacial and Holocene vegetation of the Bieszczady l\Its. (Polish Eastern Carpathians). Inst. Bot. PAN, Warszawa-Krak6w. Starkel L. 1960. Rozw6j rzezby Karpat fliszowych w holocenie. Prace Geogr. IG PAN, 22: 1-239. 1972. Charakterystyka rzezby polskich Karpat i jej znaczenie dla gospodarki ludzkiej •. Komitet Zagosp. Ziem G6rskich PAN, 10: 71-148. 1977. Paleogeografia holocenu. PWN, Warszawa. Stockmarr J. 1971. Tablets with spores used in absolute pollen analysis. Pollen et Spores· 13 (4): 615-621. Szafer W. & Jaron B. 1935. Plejstoceriskie jezioro pod Jaslem (summary: Pleistocene Lake near Jaslo in Poland). Starunia, 1: 1-20. Szafer W. 1948. P6foy glacjal w Roztokach pod Jaslem (summary: Late - Glacial in Roztoki near Jaslo - West Carpathian Mountains). Starunia, 26: 1-29. Szczepanek K. 1982. Development of the peat-bog at Slopiec and the vegetational history of the Swi~tokrzyskie (Holy Cross) Mts. in the last 10000 years. Acta Palaeobot., 22 (1): 117-130. .Sulimirski T. 1955. Polska przedhistoryczna. Cz. 1. Gryf Printers, London. - 1957-1959. Polska przedhistoryczna. Cz. 2. Gryf Printers, London. Srodon A. 1967. Swierk pospolity w czwartorz~dzie Polski (summary: The common spruce in the Quaternary of Poland). Acta Palaeobot., 8 (2): 1-59. ~ 1985. Fagus in the Forest History of Poland. Acta Palaeobot., 25 (1, 2): 119-137. Swidzinski H. 1953. Karpaty fliszowe mi~dzy Dunajcem a Sanem. Reg. Geol. Polski. Karpaty. Tektonika, 1 (2): 362-422. Swi~s F. 1980. Zarys por6wnawczej geobotanicznej charakterystyki Beskidu Niskiego z Biesz czadami i Beskidem S~deckim. Ann. Univ. Mariae Curie-Sklodowska, sec. C, 35 (8): 77-103. 1982. Charakterystyka geobotaniczna las6w Beskidu Niskiego. Analiza i synteza (summary: Geo botanical characterization of the Lower Beskid forests. Analysis and synthesis). UMCS Wydz. Biol. i Nauk o Ziemi, Lublin. Troels-Smith J. 1955. Characterization of unconsolidated sediments. Danm. Geol. Unders., IV, 3 (10): 1-73. Wi~ckowski S. & Szczepanek K. 1963. Assimilatory pigments from subfossil fir needles (Abies alba Mid.). Acta Soc. Bot. Pol., 32: 101-111. Zaki A. 1955. Poczlltki osadnictwa w Karpatach Polskich. Wierchy, 24: 99-116. Jt.. Szczepanek Acta Palaeobotanica 27/l 1 Cl ~ liJ ~ al <( l ~Ii liJ ~ l ~ 1i IO m 1: rn (/) liJ w :? w CJ) (/) 0 ~ ~ ~ l :::> E ), 0 :::> ~ ~ g ~ § E 0 ' ~ ~ :::> i I ::i ~ ~~ z I U) 0 a CJ) (/) 0 z ~ 2 ~ N :5 ~ 0: 0: ~ 0 (/) s:. x ~ ~ :> :> -~ ii: fil ~ ~ a <(I c c ~ ~ 0 ~ ffi s 0: - ~ <( Ir ' .! ~ z liJ z 3 ~ 0 :> .J 0: ::J ~ ..,:::> Q.- <( ~ ~ Ii 0 5 z ii: m :::> 0 a i= LL AP NAP IL Q. ~ ~ ~ ~~ i~ ~~ ~ CJ 0 iu oi uH ~ ~ ! ~ u 111 ~ ~ s rn ~I~ IL I ao II u ... '.-:>,'I ' ' ft I I I I• f o It I• I• I• Io Jo I Io t I t I 1 Io Io Io It It to I Io I I I I I I Io I• Io Io It It I 250 Fig. 2. Percentage pollen diagram from Jasiel mire (1981). Suma of trees, shrubs, dwarf shrubs and herbs pollen (P = 100%), excluding poilen of aquatics 'and swamp plants-and spOTes, is a calculation' base. The percentage values of excluded pollens have been calcula- ted from P + taxon. Stratigraphic symbols according to 'froels-Smith (1955) __ , ___ percentage values of taxa multiplied by 10. -x -x -x values of general pollen frequency reduced 100 times K. Seceepane1r, A.eta Palaeobotanica 27/1 j! a. i E l&I I I I IO c( II) l&I :5 I I I (I) CD :!I ~ 0 ::> ~ 1 ~ a ~ 11 ti .J (I) (I) 3 0 (I) z ~ ID N i .0 (I) ii: ~ :J :J :J :J a: c( :J l&I ~ c c I E z ... z ~ Ill l9 0: IL ~ .! :::J l&I ~ ... ~ 0 ::> J c( J ii: (I) c( j:: ,~ ~ z m :J 0 0 ~ 0 ~ l9 I iL ~ ~ 2 I m m JM 8 171 ,.,112 ,.,1111 -,., -117 7a -"' - 6 - -.., 5+- 4 3 .,., -1M --... 2 -117 200 40 - 46" Fig. 3. Pollen concentration diagram from Jasiel mire (1981). ------values of pollen concentration reduced 10 times. -x-x- vaiue$ of general pollen frequency reduced 100 times Ji, /jecsepaneJ(, Aeta. Pa.la.eobotaniea 27/l m w ! l~ ~ <( w I 1~ :J w <( w ::!i <( ~ Ill Cf!.I w <( I <( ~ ~~ _o ~ i I ~ G> i :J :J ~ ~ ::> E w -· 0: ~ (/) :::> :J 0 0 ,!i I2 ~ z Ill E >. :JI <( ~ z !:'.! 0 z ~i~ ::> Ci 2 0 ... (/) 0: w ~ c ~ G> (/) :::> LIJ n: ~ x ~ w Qz ~ ii: ~ j .c .0 :::> w D.. ~ ll. .!! 0. E C!J 0: ~ ~ :J ~ 0 ~ z iii O•:JIwz er ~1a.. > 0: ~~ (/) :J D.. ~ ~ NAP 0 C!J <( III- ii: ~o ~~ ~ ~ i c!i 5 z a: ·~!'l~lw AP l ~8 u 30 h 30 10 30 10 80 0 , ~r ~a:>~lot.;° ~·~~9?, 1 ,,,,M .,,,,1 1111111111111111111111~ 1 11 Lt>----:=]/,,__}__ _.____, ~,,,,_}_--11-1--f~ ·j~ ==-=--=--=---=-~---~~t---~~~~~={'-r-'--~-:_~~~~-::_:_-;_-;:._:_7H-n-t--1-trl-t-.,.--...j,-il--i-i.-t--+--lr--..;,_il---'--'--.--+-_ ----1 !-<::-~--:_-:,-:_ ___,_ __, 5 1--- ~ \ f-- j_ -----i 1------i 1---a.-1----1 I 1-tt11 -~.:==oo:=3 ------~1------7'/ 11---+-l----+-l-ff.-D--+-ll--,l--l---l-~~--JH rl----+---+11------l 10 ··-+-~--·--ll-4---11-4--+-+--1-4-t---l-+---l--+---ll---+---ll------l 15 "'""::.:_-;______, ~------j ~ r------~ ~ ~4- · ~-~-=--·-~-=---=-~:~ ===:=:=_=_=_=_-=_-=_-=_-=_-=_-_,1-1.J:======~::~~ ~ ~'----u..~~~-+-+--1-4-1---1-+---lf---! "-·~-~-----'! t_--~ r-1 }------1 :=:=:= ~ ~ --- ~t ==ie----!'----+-H---+-1 ~7==:=_,------i 1 ~ ~ l-"--i~------1 ""'----e \ 'J!--1- Lj . -~1----+----+---+--l-+-t ~g , ------J 1---+--J, ! L ...... ______. ~ ~ 1\--r +------4--l--1--U---1----~---+--+-----+---~----=-:=:t--1»--i--++--l--lgE_ ~~ ---d ? pr I _____t= "511>--.__------11--+--1--1-----1"~---+--1--1--+------1---'--l--~ ~'""---·· ------{~1------\1 }_ \ r 1 7 L ___JEt=1_JJ~:jt=~~======1j~11~==~-l~t/::::::~t:=:t-t:~t==== - =--=E=1Er-1if--:+tl-~-_j'l~l-~~-,il--l-co-i=--'-i=-1-· =--:_-_ -_ -_- _-_-_-_-_-'::~~~~-?~~__----lLI~__JJ_~::_-_-_-;_/:~~::~":::~:_ ..:=:~=~_,:=:~:,_-~:_ ..:=_-~::~=--~=~-~=--~=--~=-_,: Fig. 4. Percentage pollen diagram from Jasiel mire (1972). Percentag values of trees, shrubs and dwarf shrubs have been calculated from the sum of their pollen (AP). Calculation .. baso for NAP and tphagnum is the sum of AP and NAP (exluding Sphagnum)